Morphological phylogenetic analyses suggest that scalidophorans (priapulids, loriciferans, and kinorhynchs) and nematoids (nematodes and nematomorphs) form the ecdysozoan clade Cycloneuralia, which is a sister group to panarthropods. It has been proposed that extant priapulids and Cambrian priapulid-like scalidophorans, because of their conserved evolution, have the potential to illuminate the ancestral morphology, ecology, and developmental biology of highly derived ecdysozoans such as nematods and arthropods. As such, Cambrian fossils, particularly Markuelia and possibly olivooids, can inform the early evolution of scalidophorans, cycloneuralians, and ecdysozoans. However, the scalidophoran Markuelia is known exclusively as embryo fossils, and the olivooids have been alternatively interpreted as cnidarians or cycloneuralians. Here, we describe a post-embryonic scalidophoran fossil Eopriapulites sphinx new genus and species, which represents the oldest known scalidophoran, from the early Cambrian Period (∼535 Ma) in South China. E. sphinx is similar to modern scalidophorans in having an introvert armed with hollow scalids, a collar with coronal scalids, and a pharynx with pharyngeal teeth, but its scalids and pharyngeal teeth are arranged in a hexaradial pattern. Phylogenetically resolved as a stem-group scalidophoran, E. sphinx shares a hexaradial pattern with the hexaradial arrangement of certain anatomical structures in kinorhynchs, loriciferans, nematoids, and Cambrian fossils such as Eolympia pediculata, which could also be a scalidophoran. Thus, the bodyplan of ancestral cycloneuralians may have had a component of hexaradial symmetry (i.e., some but not necessarily all anatomical parts are hexaradially arranged). If panarthropods are nested within paraphyletic cycloneuralians, as several molecular phylogenetic analyses suggest, the ancestral ecdysozoans may have been a legless worm possibly with a component of hexaradial symmetry.

Morphology-based phylogenetic analyses support the monophyly of the Scalidophora (Kinorhyncha, Loricifera, Priapulida) and Nematoida (Nematoda, Nematomorpha), together constituting the monophyletic Cycloneuralia that is the sister group of the Panarthropoda. Kinorhynchs are unique among living cycloneuralians in having a segmented body with repeated cuticular plates, longitudinal muscles, dorsoventral muscles, and ganglia. Molecular clock estimates suggest that kinorhynchs may have diverged in the Ediacaran Period. Remarkably, no kinorhynch fossils have been discovered, in sharp contrast to priapulids and loriciferans that are represented by numerous Cambrian fossils. Here we describe several early Cambrian (~535 million years old) kinorhynch-like fossils, including the new species Eokinorhynchus rarus and two unnamed but related forms. E. rarus has characteristic scalidophoran features, including an introvert with pentaradially arranged hollow scalids. Its trunk bears at least 20 annuli each consisting of numerous small rectangular plates, and is armored with five pairs of large and bilaterally placed sclerites. Its trunk annuli are reminiscent of the epidermis segments of kinorhynchs. A phylogenetic analysis resolves E. rarus as a stem-group kinorhynch. Thus, the fossil record confirms that all three scalidophoran phyla diverged no later than the Cambrian Period.

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Animals with radial symmetry are abundant in the Cambrian Fortunian Stage of South China, but with relatively low diversity: representatives include Olivooides, Quadrapyrgites, carinachitiids, hexangulaconulariids and Pseudooides. Here, we report a new radial animal, Qinscyphus necopinus gen. et sp. nov., from the Fortunian small shelly fauna of southern Shaanxi Province, South China. Qinscyphus necopinus has a cup-shaped profile, with slightly raised annuli and five groups of triangular thickenings in pentaradial symmetry. This organism has a comparable morphology to, and thus a close affinity with, Olivooides and Quadrapyrgites, and is interpreted as a coronate scyphozoan. This discovery adds a new crown-group cnidarian to the Cambrian Explosion.

Although molecular studies estimated that cycloneuralians might have originated in the Ediacaran Period and diversified in the early Cambrian, it was not until 2014 that they were first reported from the Cambrian Fortunian Stage. To date only two species and five indeterminate forms have been described. Here, we report new three‐dimensionally phosphatized microfossils of cycloneuralians from the Fortunian Zhangjiagou Lagerstätte, South China. Two new genera and species, Qinscolex spinosus gen. et sp. nov. and Shanscolex decorus gen. et sp. nov., and Indeterminate Forms 1–7 are described and tentatively assigned to total‐group Scalidophora, greatly increasing the diversity of the Fortunian cycloneuralians. These Fortunian cycloneuralians have diverse cuticular spines, plates, and spinose sclerites of various sizes on the trunk, thus assignment of co‐occurring fragmented trunk parts or disassociated spines or sclerites to a specific species is currently impossible. Some of the Fortunian cycloneuralians might have reached a centimetre in length; they may have been temporary meiobenthos in their juvenile or younger adult stages, only joining the macrobenthos when they reached macroscopic dimensions. It is proposed that an introvert with hexaradially symmetric and internally hollow scalids might be the plesiomorphic state for cycloneuralians.

The latest discovery of microfossils from the lower Cambrian (Fortunian Stage) Zhangjiagou Lagerstätte in South China are presented. This lagerstätte is rich in exceptionally preserved microfossils, including embryos of Olivooides multisulcatus, Olivooides mirabilis, and Pseudooides prima; hatched stages of O. multisulcatus, O. mirabilis, Hexaconularia sichuanensis, and Quadrapyrgites quadratacris; and cycloneuralians represented by Eopriapulites sphinx. The largest known fragment of O. mirabilis implies that its adult length can be more than 9.0 mm with at least 50 annuli, and the longest known specimen of Q. quadratacris has at least 18 annuli. These unusually large specimens refute the non-feeding larvae hypothesis for Olivooides and Quadrapyrgites.Based on the current material, it is inferred that (1) early cnidarians have a high diversity in the Fortunian Stage; (2) P. prima might represent the embryonic stages of H. sichuanensis; (3) adults of Olivooides and Quadrapyrgites may have reached centimeter-scale dimensions with more than 50 annuli; (4) Olivooides and Quadrapyrgites may be better interpreted as coronate scyphozoans; (5) cycloneuralians also had a high diversity in the Zhangjiagou Lagerstätte; and (6) cycloneuralians might have originally been part of the early Cambrian meiofauna rather than belonging to the macrobenthos. Such ancestral cycloneuralians might have been Eopriapulites-like, possessing pentaradially symmetric, backward pointing, and internally hollow introvert scalids used as locomotory devices.

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